Puccinellia angustata (R. Br.) Rand and Redfield
Fl. of Mount Desert Island, Maine, 181. 1894. Poa angustata R. Br., Chlor. Melvill. 29. 1823.
Glyceria angustata (R. Br.) Fr., Nov. Fl. Suec. Mant. 3: 176. 1843. Phippsia angustata (R. Br.) A. Löve and D. Löve, Bot. Not. 128: 498. 1975 [1976].
Type: Canada. Melville Island, 1819–1820, Mr. (J.) Edwards on W. E. Parry's first voyage. (Holotype: BM! Isotype: CAN!).
Plants caespitose; densely so, culms stout, erect or prostrate, bearing only 1 to 2 leaves; less than 15 cm high, or more than 15 cm high; 10–35 cm high. Ground-level or under-ground stems not developed horizontally or vertically at, on, or below, the ground. Aerial stems erect, or decumbent; glabrous. Leaves mostly in a basal tuft. Prophylls 4–12 mm long; with scabrous veins (delicate trichomes scabrescent to glabrescent); lacking pronounced keels (but with prominent veins). Sheaths with the margins fused only in the lower part; glabrous. Ligules (0.8–)1–3 mm long (to 4 mm, Sørensen, 1953); membranous; glabrous; ovate-oblong, or transversely oblong (occasionally); apices acuminate, or acute, or obtuse, or truncate; entire. Blades 14–160 mm long; 0.5–1.2 mm wide (when rolled); appressed to the stem, or spreading; rolled in bud; without auricles (ligules decurrent); flat, or involute; midvein similar in size to other veins in the leaf. Blades adaxial surface glabrous (observed to be nearly glabrous with a few scabrous prickles, Sørensen, 1955). Blades abaxial surface glabrous.
Flowering culm nodes not exposed (usually), or becoming exposed; number visible 0–1. Flag leaf sheaths inflated, or not inflated (sheath longer than the blade). Inflorescence paniculate; dense, or diffuse; lanceolate (erect); (4–)5–8.5(–10) cm long; 4–9(–20) mm wide. Inflorescence. Inflorescence main axis glabrous, or scabrous (sparsely). Number of inflorescence branches at lowest node 2–5 (often one with a solitary spikelet). Inflorescence primary branches 6–60 mm long; scabrous (sometimes sparsely so); with appressed secondary branches (usually), or with spreading secondary branches. Spikelets. Spikelets pedicellate; disarticulating above the glumes; laterally compressed; ovate; (4–)5–8.5 mm long; 1.1–2.9 mm wide. Florets per spikelet 3–4(–6). Glumes. First glume 0.5–0.85 × the length of the second glume; 0.22–0.4 × spikelet length; 1.5–2.8 mm long; lanceolate, or ovate; glabrous; margins scabrous (appearing minutely fringed under high magnification); veins 1(–2); apex acute. Second glume 0.4 × as long as the spikelet or less; shorter than the lowest floret, or almost as long as, or longer than, the lowest floret; oblong, or elliptic; 2.2–4.2 mm long; glabrous, or with trichomes (appearing minutely fringed under high magnification). Second glume margins scabrous. Second glume veins 3 (2 of them indistinct). Rachilla internode 0.8–1.5 mm long. Rachilla 0.06–0.25 mm wide. Rachilla internode glabrous. Rachilla not pronounced between the florets; extending beyond the uppermost floret. Callus differentiated; hairs 0.1–0.5 mm long. Lemmas. Lemma 3.4–5.2 mm long; ovate, or lanceolate; rounded on the back (margins straight, thinner in texture than the body of the lemma); lemma not strongly incurved; surface dull; surface hairy; surface with trichomes on and between the veins (copiously pilose in their lower half of the veins, interveins finely pilose at base); veins 4–5. Lemma apex acute, or rounded; entire, or erose; glabrous (very rarely), or scabrous (fringed with small acute trichomes less than 25 micrometers in length); awnless. Palea well developed; 2.5–4.8 mm long; with hairy veins. Perianth reduced to lodicules. Anthers (0.6–)0.8–1.1(–1.5) mm long. Styles 2. Fruit sessile. Fruit dry; indehiscent. Fruit 1.6–2.4 mm long. Seeds 1.
Chromosome information. 2n = 42 (Bowden 1961, 5 counts; Löve and Löve 1975, 12 counts).
Distribution. Circumpolar. High arctic. Range in the Canadian Arctic Archipelago widespread. Arctic Islands: Baffin, Devon, Ellesmere, Axel Heiberg, Parry Islands, Cornwallis, Banks, Victoria.
Ecology and habitat. Substrates: river terraces, tundra (e.g., goose meadows), slopes (e.g., colonising landslides), ridges, cliffs (sometimes found near margins of glacier moraines), seashore (occasionally, on fiords); imperfectly drained moist areas, dry, moderately well drained; calcareous (on carbonates), or halophytic (occasionally found at the coast (Tanquary Fiord, 1999), and quite abundant near perennial salt springs at Expedition Fiord, Axel Heiberg Island)), or nitrophilous (Thule sites or under bird cliffs); rocks (on weathered shales, CAN 407205), gravel, sand, silt, clay, till.
Taxon as an environmental indicator. As an early coloniser this taxon is often indicative of unstable or recently disturbed environments such as land slides, clay deposits near the mouth of a river, or gravel in a river flood plain. Larger plants may be indicative of additional nutrients reaching the habitat from anthropogenic disturbances near buildings or animal activities around owl perches or lemming burrows. Plants also grow large in loose sand, to avoid being buried.
This species is reported to be non-littoral Porsild (1957, 1964), and Sorensen (1955) reports that it is probably not even halophilous. However, some specimens that clearly belong to this species have been collected by the authors along the coast on Ellesmere Island, e.g., at Tanquary Fiord. Since Puccinellia specimens of many species brought from the arctic and watered with fresh water only in greenhouses at Agriculture Canada do very well, most species may be salt tolerant but not obligate halophiles. Specimens of P. angustata also flower profusely in greenhouse conditions (personal observations, 1999–2000).
Notes. This species usually grows with a somewhat erect habit, with exterior culms generally decumbent, in clay or silt environments. In loose, windblown, riverbank sand, such as found along the MacDonald River Delta near Tanquary Fiord on Ellesmere Island, plants were found that were almost completely erect, possibly as a response to being slowly buried by sand. This is probably the most commonly collected species on the Arctic Islands, certainly so in the CAN records. Sørensen (1953) indicates that although variation within the taxon is considerable, it does not justify recognition of more than one entity in P. angustata.
The character states that have been used to key out this species in the past have differed depending on the key. For example, Sørensen (1953) indicates it has erose-ciliolate lemmas, Porsild and Cody (1980) says the lemmas are not erose-ciliolate. Whether or not this was an error or a deliberate reassessment of character states is unknown. Consaul and Gillespie (2001) found that P. angustata does have small, acuminate trichomes on the margins of the glumes and lemmas, which render the margins scabrous (see image library).
Sørensen (1955) said that his new species P. bruggemannii may easily be mistaken for stunted P. angustata, but that they are not at all alike based on leaf epidermal characters. Consaul and Gillespie (2001) found that specimens of P. bruggemannii can be distinguished from P. angustata most of the time, mostly by size characters. However, they do not agree with Sørensen’s (1955) statement that they are not at all alike. The evidence he used to support this statement, the absence of stomata on the underside of the leaf surface of P. bruggemannii, a feature that is also found only very rarely in P. phryganodes, is puzzling. Sørensen (1953, pp. 167–168) illustrates that P. phryganodes is variable in this character, with several morphotypes (including the one found in CAA) having a few scattered stomata on the leaf undersurface. The position and density of stomata on these morphotypes appears to be very similar to that of P. angustata. Therefore, his particular argument for recognizing P. bruggemannii may need more support. Consaul and Gillespie allowed that examining phenotypic plasticity in leaf epidermal cells was beyond the scope of their study, and that examining plants grown together in controlled conditions would be very useful in future studies of these two species. If the two species are only separable on size characters that are influenced by environmental factors, then P. bruggemannii may be only a depauperate form of P. angustata.
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Illustrations. • Three grasses at Eureka. Nunavut, Ellesmere island, Eureka weather station. Three grasses growing close together on clay substrate near stream. Poa hartzii near knife, Poa abbreviata (top) and Puccinellia angustata (bottom). Photograph Aug. 1991. • Plant habitat. Habitat. Nunavut, Ellesmere Island, Hazen Camp, N shore of Lake Hazen, Ellesmere Island National Park Reserve, 81°49'N, 71°20'W. Blister Hill, SW facing slope beside tiny meandering runoff stream. Silt with small rocks and pebbles, and with a yellow/white deposit at surface. 23 July 1999. L.L. Consaul 2176 & L.J. Gillespie. CAN. • Plant habitat. Plants growing on dry calcareous sandy silt. Nunavut, Ellesmere Island, Franklin Pierce Bay 79°26'N, 75°37'W. Aiken 98–029. Photograph by Mollie MacCormac. • Plant habitat. Plants growing on dry calcareous sandy silt. Nunavut, Ellesmere Island, Franklin Pierce Bay 79°26'N, 75°37'W. Aiken 98–029. Photograph by Mollie MacCormac. • Close-up of plant. Plant growing in silt with small rocks and pebbles, and with a yellow/white deposit at surface. Inflorescence diffuse. Nunavut, Ellesmere Island, Hazen Camp, N shore of Lake Hazen, Ellesmere Island National Park Reserve, 81°49'N, 71°20'W. Blister Hill, SW facing slope beside tiny meandering runoff stream. 23 July 1999. L.L. Consaul 2176 & L.J. Gillespie. CAN. • Close-up of plant. Close-up of large plant growing in loose pebble-sand of a river bank. Nunavut, Ellesmere Island, Tanquary Camp, Ellesmere Island National Park Reserve, 81°24'N, 76°52'W. S shore of MacDonald River Delta, 1.5 km NE of camp. 21 July 1999. L.L. Consaul 2154 & L.J. Gillespie. CAN. Scale bar in cm. • Plant habit. Plants growing on dry calcareous sandy silt. Nunavut, Ellesmere Island, Franklin Pierce Bay 79°26'N, 75°37'W. Aiken 98–029. Photograph by Mollie MacCormac. • Close-up of inflorescence. Close-up of pre-anthesis inflorescence, Nunavut, Ellesmere Island, Tanquary Camp, Ellesmere Island National Park Reserve, 81°24'N, 76°52'W. Pebble-sand bank on S shore of MacDonald River Delta, 1.5 km NE of camp. 21 July 1999. L.L. Consaul 2154 & L.J. Gillespie. CAN. • Close-up of inflorescence. Inflorescese pre-anthesis with closely appressed spikelets. Nunavut, Ellesmere Island, near John Richardson Bay, 80°21'N, 71°21'W. Aiken 98–053. Photograph by Mollie MacCormac. • Spikelet drawing. Puccinellia angustata distinguished from P. bruggemanni by lemma margins straight and thinner in texture than the body, not as inrolled (Sorensen 1955). Drawing reproduced from Sorensen (1952), Meddeleser om Grønland, with permission. • Close-up of lemma apex. Lemma apex as seen under 100 X, showing small marginal trichomes (less than 25 micrometers long) that are unevenly spaced along the edge and result in a sparsely scaberulous margin. Part of the apex on the left is folded down, but trichomes can still be seen. Specimen: Axel Heiberg Island, Expedition Fiord, Gillespie & Vogel 6112. (CAN). • Holotype specimen. Melville Island, 1819–1820, Mr. J. Edwards on W.E. Parry's First Voyage. Other text on label: "Puccinellia angustata (R.Br.) Nash. Poa angustata R.Br. in Parry Journ. Voy. Discov. NW Pass. Suppl. 28 (1824)!". (Holotype: BM). • Distribution map.
Cite this publication as: ‘S.G. Aiken, L.L. Consaul, and M.J. Dallwitz. 1995 onwards. Poaceae of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 10th December 2001. http://www.mun.ca/biology/delta/arcticf/’. Dallwitz (1980), Dallwitz, Paine and Zurcher (1993, 1995, 2000) , and Aiken, Dallwitz et al. (1999) should also be cited (see References).
